Diversity
Approximately, 12,000 species of ants (
Formicidae
) are documented worldwide; however, scientists estimate that closer to 15,000 actually
exist. Within this family, there are about 21 subfamilies and 283 genera. This family
of insects is highly diverse. Ants can be found on almost every continent, and they
live in various habitats, including grasslands, tropical forests, and human-dominated
areas. While they all have the same basic body plan, there are slight variations in
their appearance. Coloration ranges from black-brown to red-yellow and size may vary
depending on the species. Some species have stingers, enabling them to inject venom
into threats, while others spray formic acid as a defense mechanism. Despite variations
in geographic location and appearance, all ants are social, colonial insects, meaning
they live in large groups with each individual contributing to the collective wellbeing
of the colony. They also all have the same basic hierarchical caste system: queens,
males, and workers. Moreover, ants have incredible communication and perception capabilities.
They can find prey using powerful compound eyes, feel their environment with their
antennae, gather pheromone information with their maxillary palps and antennae, and
use their abdomens and mandibles to strike surfaces within a nest, causing vibration
messages to be sent to other ants. Lastly, ants have a wide variety of ecosystem roles
and affect humans in countless ways. They disperse seeds, aerate soil, serve as a
food source for other species, and eat decaying organic matter, thus cleaning the
environment. They even eat other household pests, help increase crop yield, and serve
as a tool for medical research. Alternatively, ants can contaminate food, harbor pathogens
dangerous to humans on rare occasions, and be major household pests, causing wood
and surface damage. Overall, ants are an interesting and diverse group of organisms,
important to their ecosystems.
Geographic Range
Ants are found on almost every continent and in most biogeographic regions, including
the Nearctic region, the Palearctic region, the Neotropical region, the Oriental region,
the Australian region, and the Ethiopian region. While many native species live in
these areas, invasive species have also been transported to these regions via human
activity.
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
- neotropical
- australian
- Other Geographic Terms
- cosmopolitan
Habitat
Ants inhabit a diverse array of terrestrial habitats. They can be found in urban areas
and suburban areas, human-dominated areas, temperate forests, tropical forests, and
grasslands. Their nests can be built in various places, including underground, in
branches, in dead trees, in human-made materials and buildings, in acorns, in leaf
litters, under stone, and so on.
- Habitat Regions
- temperate
- tropical
- terrestrial
- Terrestrial Biomes
- savanna or grassland
- forest
- rainforest
Systematic and Taxonomic History
All ants belong to the family
Formicidae
. Their distinguishing characteristic is the metapleural gland, a gland with antimicrobial
capabilities. This feature was also found in Cretaceous period fossils. This family
is related to the order
Hymenoptera
, which includes ants, wasps, and bees. They are their closest relatives. This order
is a part of the larger class
Insecta
, which includes all insects. They are characterized by having 6 legs. This class
falls under the phylum
Arthopoda
, which includes insects, centipedes, millipedes, spiders, scorpions, and crustaceans.
They are characterized by having jointed legs and an exoskeleton.
Physical Description
Generally, adult ants have segmented bodies with the following parts: head, thorax,
and abdomen. They have one pair of antennae with a bent elbow-like shape, three pairs
of walking legs, and a set of mandibles. Coloration ranges from black and brown to
yellow and red. Most species have a pair of compound eyes. Ants also have an exoskeleton
made up of a durable material called chitin for protection against the environment
and as a source of muscle attachment. Furthermore, ants are ectothermic, meaning they
use environmental heat to maintain their body temperature in an optimal range, and
they are bilaterally symmetrical, meaning that the organism can be split into two
identical halves. Sexual dimorphism exists among ants, and they also vary in appearance
depending on their role in the colony. Males are born with wings. They typically have
smaller heads but larger eyes and thorax. Females can be larger or smaller than males
depending on their designation. Queens are larger than males and are born with wings;
however, wings fall off after mating. Workers, which are all female, are smaller than
males with no wings. In some species, workers have stingers located at the bottom
of their abdomen, which inject venom into another organism. Moreover, young ants look
significantly different from adults. Eggs are typically white in color and have an
oblong shape. As larvae, ants have a worm-like appearance. Finally, pupa cacoons have
an oblong shape and are much larger than the eggs.
- Other Physical Features
- ectothermic
- bilateral symmetry
- venomous
- Sexual Dimorphism
- female larger
- male larger
- sexes shaped differently
Development
Ants undergo complete metamorphosis, meaning that the organism goes through a distinct
set of developmental stages. The four stages are the following: egg, larva, pupa,
and adult. First, ants begin as eggs that are laid by the queen. They are usually
white in color, oval-shaped, and soft. Next, the eggs hatch into larvae. Larvae are
light-colored and worm-like in shape with no legs or eyes. They must rely on adult
ants to feed and care for them. During this stage, they grow rapidly. After they reach
a large enough size, the larvae become pupae. The larvae spin a cacoon around themselves
and undergo a final transformation. The pupae look more adultlike except their limbs
are folded close to their bodies. After pupation, ants emerge as adults. Depending
on the caste of the adult, an ant will then fulfill specific duties for the colony.
The entire cycle takes a few months to complete depending on the species.
- Development - Life Cycle
- metamorphosis
Reproduction
Ants are eusocial insects. Only certain adult females can breed with males within
a colony, colony adults care for the young together, and multiple generations coexist
together. For ants, one or several queens take on the reproductive role. Queens and
males take to the air to find mates during a period known as the mating flight. Queens
then mate in the air with at least one male. Sometimes, a queen will mate inside the
nest she was born in. The male then dies within two weeks of copulation, his role
being fulfilled. After this initial mating, a queen ant never has to mate again. Her
sole purpose is to lay eggs for the colony.
- Mating System
- eusocial
Ants reproduce sexually and are gonochoric. Respectively, this means that genetic
material from two individuals, sperm and an egg, is combined to reproduce, and there
are two different sexes, male and female. Specifically, a queen ant mates with at
least one male in the air for fertilization to occur during a period called the mating
flight. In some species, however, the queen will mate inside her birth nest. Post-mating,
the male dies within two weeks. In contrast, the queen loses her wings and finds a
suitable location to start a nest and lay her eggs for the rest of her life. She never
mates again. The eggs hatch into larvae, which the queen will initially feed with
her saliva. After a certain point, enough workers will be produced. They will take
over the role of feeding and caring for the young and the queen. The queen then focuses
on laying eggs, using stored sperm from the single mating period she participated
in to fertilize them. After several years, reproductive broods are produced in a colony.
Males develop from unfertilized eggs. New queens develop from eggs fertilized by stored
sperm. These new males and queens will mate and leave the original nest to start a
new colony. Thus, the mating season for ants begins when reproductives mature and
ends with their mating exodus. The timing of the event depends on the ant species,
the climate in which they inhabit, and the weather conditions.
- Key Reproductive Features
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
- sperm-storing
- delayed fertilization
Ants dedicate a significant amount of time to parental investment for offspring. Initially,
a queen will nurture the first generation of offspring for a new colony. She will
protect and feed larvae with saliva until they pupate. They eventually emerge as adult
workers. These nonreproducing females then take over the role of feeding, caring for,
and protecting the brood. Workers travel outside the nest to find food and bring it
back to the colony to feed larvae and the rest of the community. They will also groom
pupae and transport the brood to different locations within the nest.
- Parental Investment
- female parental care
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
Lifespan/Longevity
The lifespan of an ant depends on the species and caste. In the wild, queen ants can
live from several months to about 15 years. Worker ants can live between a few months
and several years. Males live up to two weeks, dying soon after mating. In captivity,
queens may live up to thirty years.
Behavior
Ants are social, colonial insects. They live in large groups within nests, each individual
having a specific role to help the overall colony thrive. These roles are determined
by their social hierarchy system called the caste system. Ants can belong to one of
three castes: queen, male, and worker. Queens are the largest ants in the colony.
She is responsible for laying eggs. Depending on the species, a colony can have one
or several queens. Males are smaller than queens. Their sole purpose is to mate with
new mature queens who leave their original nest. They die after copulation. Lastly,
workers are the nonreproductive females. They are subdivided into major and minor
workers. Major workers are the larger workers, about the same size as a male. They
focus mainly on guarding the nest; however, they also perform other tasks similar
to minor workers. Minor workers are smaller. They maintain the nest and care for both
the brood and the queen. Specific responsibilities include the following: finding
food, storing food, building the nest, grooming the queen and brood, feeding the queen
and brood, and transporting the brood. Furthermore, ants are territorial. They will
defend their nests from rival species.
- Key Behaviors
- flies
- motile
- territorial
- social
- colonial
Communication and Perception
Ants generally communicate through chemical cues and vibrations. They perceive the
world visually, tactilely, and chemically. Ants release pheromones and create a scent
trail for other ants in the colony to follow. This may be used when one ant finds
food for the colony. Ants detect pheromones and smells via their antennae and maxillary
palps, sensory organs found at the mouth. Antennae may also be used to feel the environment
around them. Mandibles and gastors, an ant's abdomen, can be used to strike surfaces
and create vibrations. Nest mates then interpret these vibrations. Also, some ant
species rely on sight to hunt prey. Thus, these types of ants have large compound
eyes to better see the world around them.
- Other Communication Modes
- pheromones
- scent marks
- vibrations
Food Habits
Typically, ants are omnivorous. They eat a wide variety of foods: honeydew, termites,
dairy products, decaying animal and plant matter, ant larvae, sweets, starches, seeds,
and other plant parts. Ants also store food. Workers may store food internally in
fat bodies and in their gasters (abdomens), which will become enlarged. They may also
store food within the nest using seeds or dried-out insect prey and place it in chambers.
Furthermore, ants exhibit trophallaxis, the exchange of fluids between individuals
within a colony. Mature ants eat liquified forms of the food they find. Frequently,
solid food is fed to the larvae to be broken down and then it is distributed to the
rest of the colony.
- Primary Diet
- omnivore
- Foraging Behavior
- stores or caches food
Predation
Ants defend themselves by biting, injecting venom via a stinger, and spraying formic
acid. Predators include birds, mammals, arthropods, amphibians, and carnivorous plants
like pitcher plants.
Ecosystem Roles
Ants have a wide variety of ecosystem roles. Many species hold keystone positions
in their habitat. Generally, ants help disperse seeds for various plants, aerate the
soil with their nests, clean the environment of decaying organic matter, and serve
as a food source for other species. Ants also participate in mutualistic relationships
with various organisms: plants like
Acacia
and
Leonardoxa africana africana
, fungi, other insects like aphids, and bacteria. They may use certain plants as homes
like
Acacia
and
Leonardoxa africana africana
, eat byproducts of certain insects like aphids, and cultivate certain fungi for food.
In return, they protect these other species. Alternatively, certain ants can be parasites
to plants like
Leonardoxa africana africana
. Furthermore, ants serve as hosts to parasites such as the fungi
Ophiocordyceps unilateralis
, which controls the behavior of hosts and eventually kills them, and other ants like
Polyergus breviceps
, a slave-making species.
- Ecosystem Impact
- disperses seeds
- soil aeration
- keystone species
- parasite
- acacia plants ( Acacia )
- fungi ( Fungi )
- bacteria ( Bacteria )
- aphids ( Aphidoidea )
- Leonardoxa africana africana
- zombie ant fungus ( Ophiocordyceps unilateralis )
- slave-raiding ant ( Polyergus breviceps )
Economic Importance for Humans: Positive
Ants have many beneficial roles for humans. They help aerate soil as they build their
colonies, making ants useful for maintaining healthy gardens and yards. They eat other
household pests like termites (
Isoptera
), and they help increase crop yield, acting as a tool for crop pest management. Ants
are even used in medical research to develop pharmaceuticals.
- Positive Impacts
- source of medicine or drug
- research and education
- controls pest population
Economic Importance for Humans: Negative
While ants have beneficial roles, many species are considered major pests in human
households and other buildings. They infiltrate and build their nests in wood and
under stone and brick, contributing to wood and surface damage. They also contaminate
food, and some species may bite and sting humans, injecting venom or acid into the
wound. Certain species, specifically pharaoh ants (
Monomorium pharaonis
), can carry pathogens like Salmonella spp. and Staphylococcus spp. that hurt humans;
however, this occurrence is not that common.
- Negative Impacts
-
injures humans
- bites or stings
- causes disease in humans
- carries human disease
- venomous
- household pest
Conservation Status
Generally, many ant species have no special conservation status like black carpenter
ants (
Camponotus pennsylvanicus
), Argentine ants (
Linepithema humile
), and pharaoh ants (
Monomorium pharaonis
) or the population is considered secure like with red imported fire ants (
Solenopsis invicta
). Some ants are listed as vulnerable, but the information is outdated and needs to
be updated. Ants in this category include the following: slave-raiding ants (
Polyergus breviceps
), silky carpenter ants (
Camponotus universitatis
), and
Anoplolepis nuptials
.
Other Comments
Ants go out of the nest in search of food to bring back to the colony. Sometimes,
what they find is larger than they are. Thus, they are able to carry at least ten
times their own body weight.
Additional Links
Contributors
Justine Negron (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- cosmopolitan
-
having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- venomous
-
an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).
- metamorphosis
-
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
- eusocial
-
the condition in which individuals in a group display each of the following three traits: cooperative care of young; some individuals in the group give up reproduction and specialize in care of young; overlap of at least two generations of life stages capable of contributing to colony labor
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- sperm-storing
-
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
- delayed fertilization
-
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
- female parental care
-
parental care is carried out by females
- motile
-
having the capacity to move from one place to another.
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- colonial
-
used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- vibrations
-
movements of a hard surface that are produced by animals as signals to others
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- stores or caches food
-
places a food item in a special place to be eaten later. Also called "hoarding"
- soil aeration
-
digs and breaks up soil so air and water can get in
- keystone species
-
a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).
- parasite
-
an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death
- drug
-
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
- causes disease in humans
-
an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).
- venomous
-
an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
References
Anderson, G. 2022. "How Long Do Ants Live" (On-line). EcoGuard Pest Management. Accessed August 01, 2023 at https://www.ecoguardpestmanagement.com/pest-resources/how-long-do-ants-live .
Anjos, D., A. Tenas, A. Viana-Junior, R. Carvalho, H. Torezan-Silingardi, K. Del-Claro, I. Perfecto. 2022. The effects of ants on pest control: a meta-analysis. Proceedings of the Royal Society B: Biological Sciences , 289 (1981): 1-11. Accessed August 01, 2023 at https://doi.org/10.1098/rspb.2022.1316 .
Araújo, J., H. Evans, R. Kepler, D. Hughes. 2018. Zombie-ant fungi across continents: 15 new species and new combinations within Ophiocordyceps. I. Myrmecophilous hirsutelloid species. Stud Mycol. , 90: 119–160. Accessed July 31, 2023 at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6002356/ .
Beatson, S. 1972. Pharaoh's ants as pathogen vectors in hospitals. The Lancet , 299 (7747): 425-427. Accessed August 01, 2023 at https://www.sciencedirect.com/science/article/pii/S0140673672908690 .
Bennett, K., A. Ellison. 2021. "Ants of the North Quabbin Region" (On-line). Harvard Forest. Accessed July 24, 2023 at https://harvardforest.fas.harvard.edu/ants/body-structure .
Blatrix, R., S. Bouamer, S. Morand, M. Selosse. 2009. Ant-plant mutualisms should be viewed as symbiotic communities. Plant Signal Behav. , 4 (6): 554–556. Accessed July 31, 2023 at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2688311/ .
Brady, S., T. Schultz, B. Fisher, P. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proc Natl Acad Sci USA , 103 (48): 18172–18177. Accessed August 02, 2023 at https://doi.org/10.1073/pnas.0605858103 .
Cetintas, R. 1998. The University of Florida Book of Insect Records . Gainesville, FL: University of Florida. Accessed August 01, 2023 at https://entnemdept.ufl.edu/walker/ufbir/contribute_copyright.shtml .
Charles, V., S. C., R. Kavitha. 2022. Diversity of ant species (Hymenoptera: Formicidae) in Teresian College campus, Mysuru, Karnataka: A model ecosystem of habitat persistence. Annals of Entomology , 40 (2): 59-61. Accessed July 22, 2023 at https://ezproxy2.library.colostate.edu/login?url=https://search.ebscohost.com/login.aspx?direct=true&AuthType=cookie,ip,url,cpid&custid=s4640792&db=asn&AN=162861795&site=ehost-live .
Chérasse, S., S. Aron. 2018. Impact of immune activation on stored sperm viability in ant queens. Proc. R. Soc. B. , 285 (1893): 1-7. Accessed August 04, 2023 at https://doi.org/10.1098/rspb.2018.2248 .
Cleveland Clinic, 2022. "Ant Bites" (On-line). Cleveland Clinic. Accessed August 01, 2023 at https://my.clevelandclinic.org/health/diseases/22943-ant-bites#:~:text=During%20an%20ant%20bite%2C%20the,stings%20can%20be%20very%20painful. .
Cornell University, 2023. "Ants" (On-line). Cornell CALS: College of Agriculture and Life Sciences. Accessed July 22, 2023 at https://cals.cornell.edu/new-york-state-integrated-pest-management/outreach-education/whats-bugging-you/ants .
Del Toro, I., K. Towle, D. Morrison, S. Pelini. 2013. Community structure and ecological and behavioral traits of ants (Hymenoptera: Formicidae) in Massachusetts open and forested habitats. Northeastern Naturalist , 20 (1): 103-114. Accessed July 22, 2023 at https://ezproxy2.library.colostate.edu/login?url=https://search.ebscohost.com/login.aspx?direct=true&AuthType=cookie,ip,url,cpid&custid=s4640792&db=asn&AN=87308725&site=ehost-live .
Drees, B., B. Summerlin. 2023. "House-infesting ants and their management" (On-line pdf). Texas Agriculture Extension Service: The Texas A&M University System. Accessed July 29, 2023 at https://liveoak.agrilife.org/files/2011/07/House_Infesting_Ants_Management_15.pdf .
Duncan, K., J. Philippoff, F. Pottenger, L. Kaupp, M. Lurie, D. Lin, E. Baumgartner. 2023. "Introduction to Phylum Arthropoda" (On-line). Exploring Our Fluid Earth: Teaching Science as Inquiry. Accessed August 02, 2023 at https://manoa.hawaii.edu/exploringourfluidearth/biological/invertebrates/phylum-arthropoda#:~:text=The%20phylum%20Arthropoda%20contains%20a,3.72). .
Fernández, F., R. Guerrero, A. Sánchez-Restrepo. 2021. Systematics and diversity of Neotropical ants. Revista Colombiana de EntomologĂa , 47 (1): e11082. Accessed August 04, 2023 at http://www.scielo.org.co/scielo.php?script=sci_arttext&pid=S0120-04882021000100011 .
Franks, N., A. Sendova-Franks. 1992. Brood sorting by ants: distributing the workload over the work-surface. Behavioral Ecology and Sociobiology , 30: 109–123. Accessed July 31, 2023 at https://link.springer.com/article/10.1007/BF00173947 .
Hahn, J. 2020. "Ants" (On-line). University of Minnesota Extension. Accessed July 28, 2023 at https://extension.umn.edu/insects-infest-homes/ants#:~:text=Most%20ants%20are%20brownish%20to,to%201%2F2%20inch%20long. .
Helms, J. 2023. Climate, geography, and the mating phenology of ants. Insectes Sociaux , 70: 119-125. Accessed July 30, 2023 at https://link.springer.com/article/10.1007/s00040-022-00888-y .
Holbrook, T. 2023. "Face to Face with Ants" (On-line). Arizona State University: Ask a Biologist. Accessed July 24, 2023 at https://askabiologist.asu.edu/explore/ant-anatomy .
Hölldobler, B. 1999. Multimodal signals in ant communication. Journal of Comparative Physiology A , 184: 129–141. Accessed July 22, 2023 at https://link.springer.com/article/10.1007/s003590050313 .
Klotz, J., J. Mangold, K. Vail, L. Davis, R. Patterson. 1995. A survey of the urban pest ants (Hymenoptera: Formicidae) of peninsular Florida. The Florida Entomologist , 78 (1): 109-118. Accessed August 01, 2023 at https://doi.org/10.2307/3495674 .
Kumari, J., R. Sah, N. Mohaideen, S. Ahmad, S. Pati, S. Singh. 2022. Studying the rationale of fire ant sting therapy usage by the tribal natives of Bastar revealed ant venom-derived peptides with promising anti-malarial activity. Toxins (Basel) , 14 (11): 789. Accessed August 01, 2023 at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9697016/ .
Lach, L., C. Parr, K. Abbott, E. Wilson. 2010. Ant Ecology . Oxford, England: Oxford University Press. Accessed July 31, 2023 at https://www.google.com/books/edition/Ant_Ecology/vlwVDAAAQBAJ?hl=en&gbpv=1 .
Leahy, L., B. Scheffers, S. Williams, A. Andersen. 2020. Diversity and distribution of the dominant ant genus Anonychomyrma (Hymenoptera: Formicidae) in the Australian wet tropics. Diversity (14242818) , 12 (12): 474. Accessed July 23, 2023 at https://ezproxy2.library.colostate.edu/login?url=https://search.ebscohost.com/login.aspx?direct=true&AuthType=cookie,ip,url,cpid&custid=s4640792&db=asn&AN=147754442&site=ehost-live .
Lubertazzi, D. 2018. Ant (Formicidae) assemblage in South Africa’s Vachellia (Acacia) karroo thorns. Transactions of the American Entomological Society (1890-) , 144 (1): 119-130. Accessed July 31, 2023 at https://www.jstor.org/stable/26450478?seq=1 .
Martins, L., J. Serrao, H. Santos, V. Araujo. 2022. Anatomy and histology of the metapleural gland in the giant tropical ant Paraponera clavata (Fabricius, 1775) (Formicidae: Paraponerinae). An Acad Bras Cienc , 94 (2): e20201368. Accessed August 04, 2023 at https://www.scielo.br/j/aabc/a/TksQ8LVKkj3sBwvj83PmgKF/?format=pdf&lang=en .
Meurville, M., A. LeBoufe. 2021. Trophallaxis: the functions and evolution of social fluid exchange in ant colonies (Hymenoptera: Formicidae). Myrmecological News , 31: 1-30. Accessed July 31, 2023 at https://www.biotaxa.org/mn/article/view/66610 .
Michelson, M. 2016. "Parasitic Ants and Their Slaves" (On-line). California Academy of Sciences. Accessed July 31, 2023 at https://www.calacademy.org/explore-science/parasitic-ants-and-their-slaves .
NC State University, 2015. "The Exoskeleton" (On-line). NC State Agricultural and Life Sciences. Accessed July 24, 2023 at https://genent.cals.ncsu.edu/bug-bytes/exoskeleton/#:~:text=The%20Exoskeleton,sensory%20interface%20with%20the%20environment. .
Santoandré, S., I. Bellocq, J. Filloy. 2016. Southernmost record and new habitat type for Eurhopalothrix bruchi (Santschi, 1922) (Hymenoptera: Formicidae) in Sierra de La Ventana (Buenos Aires, Argentina). Check List , 12 (4): 1-5. Accessed July 22, 2023 at https://ezproxy2.library.colostate.edu/login?url=https://search.ebscohost.com/login.aspx?direct=true&AuthType=cookie,ip,url,cpid&custid=s4640792&db=asn&AN=134704226&site=ehost-live .
Siddiqui, J., B. Bamisile, M. Khan, W. Islam, M. Hafeez, I. Bodlah, Y. Xu. 2021. Impact of invasive ant species on native fauna across similar habitats under global environmental changes. Environmental Science and Pollution Research , 28: 54362–54382. Accessed July 31, 2023 at https://link.springer.com/article/10.1007/s11356-021-15961-5 .
Simothy, L., F. Mahomoodally, H. Neetoo. 2018. A study on the potential of ants to act as vectors of foodborne pathogens. AIMS Microbiol. , 4 (2): 319–333. Accessed August 01, 2023 at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6604928/ .
Smith, C. 2023. Sexual dimorphism as a facilitator of worker caste evolution in ants. Ecology and Evolution , 13 (2): 1-10. Accessed July 24, 2023 at https://doi.org/10.1002/ece3.9825 .
Sorvari, J. 2022. Biogeography and habitat preferences of red wood ants of the Formica rufa group (Hymenoptera: Formicidae) in Finland, based on citizen science data. European Journal of Entomology , 119: 92-98. Accessed July 22, 2023 at https://ezproxy2.library.colostate.edu/login?url=https://search.ebscohost.com/login.aspx?direct=true&AuthType=cookie,ip,url,cpid&custid=s4640792&db=asn&AN=160158556&site=ehost-live .
The IUCN Red List, 2023. "The IUCN Red List" (On-line). The IUCN Red List of Threatened Species. Accessed August 02, 2023 at https://www.iucnredlist.org/search?query=Ants&searchType=species .
The Louisiana Department of Wildlife and Fisheries, 2023. "Red Imported Fire Ant" (On-line). The Louisiana Department of Wildlife and Fisheries. Accessed August 02, 2023 at https://www.wlf.louisiana.gov/species/detail/red-imported-fire-ant .
The National Wildlife Federation, 2023. "Ants" (On-line). The National Wildlife Federation. Accessed August 02, 2023 at https://www.nwf.org/Educational-Resources/Wildlife-Guide/Invertebrates/Ants .
University of California Agricultural and Natural Resources, 2017. "How to Manage Pests: Key to Identifying Common Household Ants" (On-line). UC IPM. Accessed July 29, 2023 at https://ipm.ucanr.edu/TOOLS/ANTKEY/antid1.html .
Wetterer, J. 2012. Worldwide spread of the African big-headed ant, Pheidole megacephala (Hymenoptera: Formicidae). Myrmecological News , 17: 51-62. Accessed July 24, 2023 at https://www.researchgate.net/publication/257747995_Worldwide_spread_of_the_African_big-headed_ant_Pheidole_megacephala_Hymenoptera_Formicidae .
Wilson, E., B. Hölldobler. 2005. Eusociality: Origin and consequences. PNAS , 102 (38): 13367-13371. Accessed July 30, 2023 at https://doi.org/10.1073/pnas.0505858102 .
Yates, J. 1992. "Ants" (On-line). Urban Knowledge Master. Accessed July 31, 2023 at http://www.extento.hawaii.edu/kbase/urban/site/ants.htm .